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70 TROP. LEPID. RES., 18(2):70-73, 2008 DANIELS et al.: Immature stages of Panacea procilla

THE BIOLOGY AND IMMATURE STAGES OF PANACEA PROCILLA LYSIMACHE (: ) FROM COSTA RICA, WITH THE REPORT OF A NEW LOCALITY RECORD

Jaret C. Daniels,1, 2 Ernesto Rodriguez,3 and J. Court Whelan1, 2 1Department of Entomology and Nematology, University of Florida, P.O. Box 110620, Gainesville, FL 32611, USA 2McGuire Center for Lepidoptera and Biodiversity, Florida Museum of Natural History, P.O. Box 112710 , Gainesville, FL 32611, USA 3El Bosque Nuevo, San Jose, Costa Rica

Abstract - Descriptions of the immature stages of Panacea procilla lysimache are presented, along with new locality records for this butterfly and its larval host in Costa Rica. Basic natural history, larval behavior, larval host use, and the distribution of Panacea procilla lysimache are discussed.

Key Words: Batesia, biology, Caryodendron, , Guanacaste, host , immature stages, larvae, Neotropical, Nicaragua, pupae.

The Neotropical Panacea Godman and Salvin occurs from enclosure. Costa Rica south through Colombia, Venezuela, the Guianas, the LARVAL HOST PLANT Amazon Basin, and into Bolivia (D’Abrera, 1987; DeVries, 1987; Emmel & Austin, 1990; Penz & DeVries, 2002). Members are Caryodendron angustifolium Standley (Euphoriaceae) (Fig. large, conspicuously marked butterflies characterized by iridescent 14-17) was utilized for adult oviposition and larval feeding. This blue-green dorsal bands and colorful ventral hind wing surfaces. rare rainforest tree appears to be known from only a few isolated Although widespread and often abundant components of tropical specimen records including western Colombia (Antioquia and forest faunas in South America, specimens are rare and poorly Chocó), Estación Biológica La Selva in Heredia, Costa Rica, and represented in collections from Central American locations; in fact, the type locale of Progresso in Chiriquí, Panama, all below 1000 the subspecies’ known distribution was solely limited to Panama m (Berger and Huft, 1995; Mitre, 1998, 1997; Taylor and Gómez, until the first record of Panacea procilla lysimache Godman 2008;Webster and Huft, 1988; Woodson, 1943). Small numbers of and Salvin from Finca la Selva, Costa Rica by DeVries in 1989. the plant occur naturally on the El Bosque Nuevo property along Similarly, detailed information on the life history, ecology and with many dozen more currently in culture. behavior of the various and subspecies remains generally limited. IMMATURE STAGES As part of a broader ecological research project dealing with the impact of butterfly farming on local species abundance, we were Egg (Fig. 5). Cream to white; approximately round, with base and micropylar able to study the natural history of a Panacea procilla lysimache region flattened; chorion with 16 longitudinal ribs, each bearing spines that increase in length from the base to the edge of the micropylar region, where they form a (Fig. 1-4) population in extreme northwestern Costa Rica. Here, crown. Egg darkens to yellow with embryonic development. we present information on the biology of the immature stages. Fifth Instar Larva (Fig. 6-7). 42-45 mm long; head black with short setae and MATERIALS AND METHODS adorned with a pair of white-tipped, black dorsolateral scoli (to approximately 15 mm in length or over three times as long as the height of head); scoli densely covered with small tubercles bearing a short seta, and 9-10 prominent spines The immature stages and general biology of Panacea procilla loosely distributed around the shaft from the base to over one half the length of lysimache were studied in the field and under laboratory the scoli; postgenal region on each side of head bearing spines. Body orange with conditions from December 2006 to October 2007 at El Bosque white prothorax; thoracic legs, spiracles and proleg shields black; body scoli black on black chalazae (except where noted otherwise); scoli long and sparsely covered Nuevo, a privately owned 180 ha field station and butterfly farm with spinules; branched body scoli with stem; T1 with white prothoracic plate; in Guanacaste Province, northwestern Costa Rica, approximately bifurcated white subdorsal scoli with black tip, a branched white supraspiracular 8 km north of Santa Cecilia and 12 km south of Lake Nicaragua scoli with black tip, and a simple subspiracular scoli with black tip; T2 with five- (11o03’30.4 N”; 85 o21’26.2” W) at altitudes between 307-355m. branched subdorsal scoli off short stem, three-branched supraspiracular scoli off short stem , and simple subspiracular scoli, cuticle in the anterior dorsal portion The research site and surrounding area can be characterized as of T2 darkened (variable) and rough; T3 with five-branched subdorsal scoli, a network of primary and secondary lower montane rainforest three-branched supraspiracular scoli, and two simple subspiracular scoli aligned interspersed with agroforesty plots, pasture, disturbed sites, homes horizontally; A1 to A7 with single unbranched dorsal scolus on a small chalaza and roads. Observations of adults were made in the immediate located on anterior portion of each segment; A7 with a single three-branched dorsal scolus with stem on a large chalaza located at the posterior end of the segment; A8 vicinity of the farm and were followed by concentrated searches with a single five-branched dorsal scolus with stem on a large chalaza located at for the immature stages in the surrounding landscape. Samples the posterior end of the segment; A2 to A7 with single three-branched subdorsal of larvae were initially reared on cut host leaves placed in small scoli with stem (A7 sometimes with four-branched scoli); A8 with five-branched plastic containers. All rearing containers were housed in a dedicated subdorsal scoli with stem; A2 to A8 with bifurcated superspiracular with short stem; A2 to A6 with three subspiracular scoli, one bifurcated off short stem, remainder rearing laboratory with screen windows and maintained under an simple; A7 and A8 with two simple subspiracular scoli (sometimes bifurcated on ambient temperature regime. Following nursery propagation of A8); A9 with five-branched subdorsal scoli,; anal plate oval, black and convex with plants from vegetative samples collected on site, induction of adult a pair of small simple scoli. oviposition and subsequent larval culture was completed using Sixth Instar Larva. 58-63 mm; our observations indicate that some larvae may occasionally go through supplementary instars. Six larval instars were documented living potted plant material maintained in a large outdoor screen periodically. Although not observed, it is possible that more than six larval instars DANIELS et al.: Immature stages of Panacea procilla DANIELS et al.: Immature stages of Panacea procilla TROP. LEPID. RES., 18(2):70-73, 2008 71

Figs. 1-4. Adults of Panacea procilla lysimache from El Bosque Nuevo property, Guanacaste Province, Costa Rica. 1, male, dorsal view. 2, male, ventral view. 3, female, dorsal view. 4, female, ventral view.

are produced. Similar to fifth in morphology except noticeably larger in size and P. procilla lysimache will soon be recorded as part of the butterfly more yellow-orange in color. fauna of Nicaragua. Pupa (Fig. 11-13). 26-38 mm long; pendant, although pupa may be suspended All wild adults were observed as isolated individuals. Courtship more horizontal to substrate. Overall cream-yellow to bright yellow with black and mating were not observed, even under captive situations. spots and lines; elongate and cylindrical in shape, and without projections. Head Eggs were laid singly on the underside of mature host leaves. with oblique dorsolateral crests; anterior portion of crest with black spot; black All larval instars were moderately gregarious, especially during spot anterior to eye; black spot on eye; surface of wing with transverse elongated black spots and eight triangular black spots along posterior margin; abdomen with molting (Fig. 10), but last instar larvae pupated solitarily. Larvae incomplete longitudinal dorsal, subdorsal, spiracular, ventrolateral, and ventral were almost always observed feeding on mature host leaves rather rows of black spots; attached by black cremaster to silk pad on substrate. than new terminal growth. Late instar larvae frequently rested on the upper surface of mature leaves along the mid-vein when DISCUSSION not actively feeding (Fig. 9). When disturbed or approached by an observer, larvae would raise their anterior segments off the Our observations at El Bosque Nuevo represent new locality substrate at a 45-90o angle and bounce fiercely (and synchronously records for both P. procilla lysimache and C. angustifolium in if multiple larvae were present). Similar behavior was observed in Costa Rica. Documentation of multiple adult individuals over pupae, which would either repeatedly bounce up and down or snap several months and of both sexes indicates the presence of a local violently from side to side. This vigorous response to disturbance breeding population in the area. Yhe new site’s elevation (307- may be effective at startling approaching predators or thwarting 355 m) is higher than that of Estación Biológica La Selva (35-130 direct attack. m) where all previous Costa Rican records originate. The close The information presented here on P. procilla lysimache, coupled proximity of El Bosque Nuevo to Lake Nicaragua suggests that with that of DeVries (2001) and Teshirogi (2007), advances our 72 TROP. LEPID. RES., 18(2):70-73, 2008 DANIELS et al.: Immature stages of Panacea procilla DANIELS et al.: Immature stages of Panacea procilla DANIELS et al.: Immature stages of Panacea procilla TROP. LEPID. RES., 18(2):70-73, 2008 73

understanding of the basic biology of the charismatic Neotropical (Lepidoptera: Nymphalidae). Tropical Lepidoptera 10:43-46. genus Panacea and helps further illustrate the morphological DeVries, P. J. and T. R. Walla 2001. Species diversity of and community structure in neotropical fruit- similarity of its immature stages and host choice to those of feeding butterflies.Biological Journal of the Linnean Society, 74:1-15. Batesia hypochlora Felder and Felder. This study also emphasizes Emmel, T. C. and G. T. Austin the collaborative research opportunities available with the growing 1990. The tropical rain forest butterfly fauna of Rondônia, Brazil: species global butterfly farming and ranching industry. diversity and conservation. Tropical Lepidoptera, 1:1-12. Mitre, M. E. 1997. Caryodendron angustifolium. In: IUCN 2007. 2007 IUCN Red List of ACKNOWLEGMENTS Threatened Species. . Penz, C. M. and P. J. DeVries We thank Mr. John Fazzini and all the staff at El Bosque Nuevo 2002. Phylogenetic analysis and review of Panacea and Batesia butterflies (Nymphalidae). Journal of the Lepidopterists’ Society, 56(4): 199-215. for their kind assistance and support. Taylor, C. and L. D. Gómez 2008. La Flórula Digital de la Estación Biológica La Selva. . Teshirogi, M. Burger, W. and M. Huft 2007. The splendid in the Neotropical region. Butterflies, 47: 30- 1995. Family 113 Euphorbiaceae. Fieldiana: Botany, New Series 36: 1–169. 44. D’Abrera, B. Webster, G. L. and M. J. Huft 1987. Butterflies of the Neotropical Region. Part III. Brassolidae, Acraeidae, 1988. Revised synopsis of Panamanian Euphorbiaceae. Annals of the Missouri Nymphalidae (partim). Hill House, Victoria. Botanical Garden, 75(3): 1087–1144. DeVries, P.J. 1987. Butterflies of Costa Rica and their Natural History. Vol. 1: Papilionidae, Pieridae, Nymphalidae. Princeton University Press, Princeton. DeVries, P. J. 1989. Notes on Panacea procilla lysimache (Nymphalidae) from Costa Rica. Journal of Research on the Lepidoptera 27: 140-141. DeVries, P. J., C. M. Penz, and T. R. Walla 1999. The biology of Batesia hypochlora in an Ecuadorian rainforest

Figs. 5-17. Immature stages and larval host plant of Panacea procilla lysamache, El Bosque Nuevo property, Guanacaste Province, Costa Rica. 5, egg, immediately following ovipostion. 6, fifth instar larva, lateral view. 7, fifth instar larva, dorsal view. 8, head capsule from fifth instar larva.9 , fifth instar larvae resting on med-vein of mature host plant leaf. 10, larvae, note gregarious nature during molting. 11. pupa, dorsal view. 12, pupa, lateral view. 13, pupa, ventral view. 14, young leaf of Caryodendron angustifolium. 15, mature leaf of Caryodendron angustifolium. 16, Caryodendron angustifolium plant in culture at El Bosque Nuevo. 17, Caryodendron angustifolium with fruit.